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Home , Acarapis, Aceria sheldoni, Aculops lycopersici, Amblyseius, Amblyseius andersoni, Amblyseius largoensis

CAB Reviews 2018 13, No. 031

The underestimated worth of predatory and parasitic in India: does it really have to import exotic for biological control?

Salil Kumar Gupta1 and Prakya Sreerama Kumar2*

Address: 1Medicinal Plants Research and Extension Centre, Ramakrishna Mission, Narendrapur, Kolkata 700 103, India. 2Division of Germplasm Conservation and Utilisation, ICAR–National Bureau of Agricultural Resources, P.O. Box 2491, H.A. Farm Post, Hebbal, Bengaluru 560 024, India.

*Correspondence: Prakya Sreerama Kumar. Email: [email protected]

Received: 22 August 2017 Accepted: 15 August 2018 doi: 10.1079/PAVSNNR201813031

The electronic version of this article is the definitive one. It is located here: http://www.cabi.org/cabreviews

© CAB International 2018 (Online ISSN 1749-8848)

Abstract

Predatory and parasitic mites are likely to assume a greater role in crop protection as world attention is now focused on finding out alternative means of agricultural management, and some may be beneficial in the medical and veterinary fields. This review on the status of research on predatory and parasitic mites in India is written in the light of the recent push for introduction of exotic phytoseiid mites into the country. It provides an overview of the predatory mites associated with agri-horticultural crops as well as of some of public health importance in India and also of some known from outside the country for appreciating their potential. It has information on predatory mites representing 27 families under three orders, viz. , (suborder ), (suborder , cohort and also cohort excluding Astigmatina). Since phytoseiid mites are the most dominant, highly effective, economically important and widespread predators, the major emphasis has been laid on this family. The promising predators that emerged out of this review are: channabasavannai, A. herbicolus, A. largoensis, alstoniae, E. finlandicus, E. ovalis, fallacis, N. longispinosus, Scapulaseius suknaensis, Transeius tetranychivorus (); Agistemus fleschneri, A. industani, Eryngiopus coimbatorensis (Stigmaeidae); baccarum, Walzia indiana (); setirostris (); fleshneri (Iolinidae); and tritici (), all associated with various agri-horticultural crops; Microtrombidium saharanpuri and species parasitizing housefly and mosquitoes, respectively. Since many of the predatory families appear to be still unexplored, those need immediate attention for exploration, documentation and profitable exploitation. The need to give more emphasis and encouragement to indigenous predatory mites than to exotic species is also discussed.

Keywords: Biological control, Importation, Natural enemies, Pest management

Review Methodology: This review has been prepared based on the information collected from published books, particularly the ones on India-specific written by the senior author, and from several online resources, including CAB Direct (www.cabdirect.org). Further, a few important textbooks and manuals on mites were also consulted, and the cross-references from the bibliography collated thus were sourced to obtain further information for the review. As far as possible, an up-to-date review has been attempted by thoroughly checking with publicly available online resources as well.

Introduction and insects, and pathogens, but not parasitoids. Similarly, several insect pests of medical and veterinary Phytophagous mites, which cause enormous losses to importance are often debilitated by parasitic mites in agriculture and horticulture, are often decimated in the nature. In view of this importance, attempts have been field by their natural enemies that include predatory mites made worldwide, including in India, to survey and

http://www.cabi.org/cabreviews 2 CAB Reviews document the predatory and parasitic mites for controlling prey upon a variety of nematodes and small [8]. agricultural pests and also those pests which are of medical This family has not received enough attention in India. and veterinary importance. The present review aims at giving a critical overview on predatory and parasitic mites to highlight how much is known and what should be the future strategy with Family Blattisociidae special reference to the Indian context so as to achieve sustainable agriculture and reduce dependency as much This is an assemblage of diverse groups that occur in as possible on chemical control measures. This review terrestrial, arboreal and subaquatic habitats [9, 10]. A total assumes special significance in the light of efforts by of 389 species of Blattisociidae are known from the world multinational pest control companies to extensively intro- [11], while India represents about 33 species. These large, duce exotic predatory mites such as Amblyseius swirskii brownish or whitish mites prey upon nematodes, micro- (Athias-Henriot) and (McGregor) arthropods and fungi [12]. into India. Risks such as non-target effects, changes in the Blattisociids are common predators of acarid mites, trophic web as well as threats to the environment and and eggs and larvae of insects in stored products and indigenous are of high concern wherever nests of small mammals and birds [13, 14]. Some are exotic species are considered for introduction. Though uninvited pests of laboratory cultures of arthropods [15]. exotic species have been released in several countries Some are phoretic and several species occur in rotten without even considering the use of native species, a logical habitats [16]. way to reduce the risks involved in releasing exotic species, Blattisocius tarsalis (Berlese) is a good predator of a particularly for augmentative biological control, would be number of tetranychid mites and . It controls stored to use native species [1]. Unfortunately, the above- grain pests under controlled conditions [17]. Adult mentioned exotic agents are meant for augmentative blattisociine females are known to attach themselves to biological control and not for classical biological control moths phoretically and may feed on the eggs and larvae of of pests in India. the hosts [18]. Though importation of predatory mites was not new to Over 150 species are known from the world under the India, the attempts were not as fruitful as anticipated. For Lasioseius [19]. Several Lasioseius species, including instance, in September 1961, newsami (Evans) Lasioseius parberlesei (Tseng), are good natural enemies of [=Scapulaseius newsami (Evans)] was imported from the panicle mite, spinki (Smiley), and Malaysia for trials against several mite species in tea, but can suppress the pest population. Lasioseius scapulatus could not be established [2]. persimilis (Kennett) is an important species for nematode control. (Athias-Henriot) was first imported by the Indian Station Lasioseius safroi (Ewing) preys upon nematodes under the of Commonwealth Institute of Biological Control in 1965 bark of pine infested with Ips beetles [16]. Lasioseius species from Chile via for use against Tetranychus are efficient predatory mites in Australia [20]. Adult females species on several crops [3]. Though it was recovered of some species are phoretic on hummingbirds [21]. from the released sites, eventually it could not be used Blattisociids associated with plants have not been ade- for the intended purpose. In 1984, P. persimilis was quately explored in India. again imported from the Glasshouse Crops Research Institute, Littlehampton in England, for research under the All-India Coordinated Research Project on Biological Control of Crop Pests and Weeds [4]. Unfortunately, Family even that attempt was not sustainable. Similarly, Amblyseius chilenensis (Dosse) [=Neoseiulus chilenensis (Dosse)], This is a diverse group both morphologically and behav- which was imported from the USA in 1984 for use against iourally. These large, brownish mites are free-living Tetranychus species [5], met with the same fate. or associated with arthropods, birds or mammals. Some are predators, some nest-dwelling and some obligatory or facultative parasites of birds and mammals [16]. They Order Mesostigmata are good predators of soil insects and nematodes, helping in biological control. Hypoaspsis aculifer Family (Canestrini) brought about control of the plant nematode Tylenchorhynchus dubius (Buetschli) in a pot experiment This family includes two subfamilies, Arctoseiinae and [22]. Haemogamasus pontiger (Berlese) feeds on acarid Ascinae. These are free-living predators that occupy mites in bird nests in [23]. The nests having this diverse habitats and prey upon nematodes, micro- mite had fewer fleas as well. Stratiolaelaps scimitus arthropods and eggs of sciarid flies [6]. Asca species are (Womersly) is known as a biocontrol agent of dipteran known to consume other mites, nematodes and small pests on mushrooms [24]. Laelapids have not received insects such as collembolans [7]. Species of Antennoseius enough attention in India.

http://www.cabi.org/cabreviews Salil Kumar Gupta and Prakya Sreerama Kumar 3 Family Macrochelidae Khot). However, this family has not been investigated well in India, and only one species as a predator has been It is an assemblage of 400 species under 15 genera from the reported [38]. world. Macrochelids are deep brownish, well-sclerotized, large mites that are mostly predaceous on nematodes and eggs/larvae of arthropods. They also occur in organic Family Phytoseiidae substrates and have been recorded from fur of mammals and occasionally on birds [25]. Macrochelids have been Phytoseiidae is probably the best worked out, well- widely used for control of dipteran pests in greenhouses explored and best utilized family of predatory mites [39, and mushroom cultivation [24]. Some macrochelids can be 40]. Consultation to any abstracting journal will reveal good predators of immatures of housefly, Musca domestica hundreds of references in a variety of fields of Phytoseiidae L., and can be considered as natural biocontrol agents. every year and that indicates what a tremendously Al-Dulaimi [26] studied rate and developmental important role the phytoseiid mites have in biocontrol time of Macrocheles glaber (Muller) on frozen housefly. programmes. Kostiainen and Hoy [41] published a bibli- According to de Azevedo et al. [27], M. glaber also preys on ography of Phytoseiidae comprising publications on the springtails, phorid flies and nematodes. It can predate on biological aspects of these mites, for a total of 4634 the Australian bush fly, Musca vetustissima (Walker), and references covering a period of 1960–1994. almost total control could be achieved through this mite. Phytoseiids are free-living, terrestrial mites that occur Interestingly, the intrinsic rate of increase, finite rate of on plant foliage, tree bark and surface, and occasionally increase, net reproductive rate and net predation rate for in litter and other habitats. Generally, these mites are M. glaber were all significantly lower than those of highly predaceous on different species of Tetranychidae, S. scimitus, a laelapid known to be a generalist predator [28]. and . They are usually whitish, Macrocheles peregrinus Krantz was introduced from creamish, light brownish or reddish and often their colour Australia to South Africa to effectively control the bush changes after consuming preys. The prey density is the fly and buffalo fly [29]. Macrocheles robustulus (Berlese) is a crucial factor that determines the response of predators. good predator of [30], and is available commercially. Thus, the success of a phytoseiid will depend upon Macrocheles autumnalis (DeGeer) is another species that functional and numerical responses as well as on prey can bring about good control of stable fly, Stomoxys colonization pattern. The rate of food intake is often related calcitrans (L.) [31]. Macrochelids have not been investigated to egg production. adequately in India. (Indian scenario vis-à-vis world scenario) Since the publication of the world review of Phytoseiidae Family Melicharidae [42], hundreds of workers have contributed from all around the world on exploration and documentation This diverse group of mites occupies terrestrial and of phytoseiid mites. Those references are numerous and above-ground habitats [9]. These mites are adapted to mentioning even a fraction here will be an impossible task. feeding upon fungi, and nectar, and there is one Moraes et al. [43], Chant and McMurtry [44], Prasad [45] species that is parasitic to cockroaches [32]. Proctolaelaps and Demite et al. [46] are some of the recent reviews giving pygmaeus (Muller) feeds upon a variety of other acarines updated information on phytoseiid taxonomy. As per the [33]. The adult females of Proctolaelaps also show phoresy Phytoseiidae Database, 2479 species are valid out of the [34]. Adult females of Rhinoseius, Tropicoseius and some 2763 described species from around the world [47]. As for Proctolaelaps show phoretic behaviour with hummingbirds, India, Gupta’s [38, 48–50] periodic reviews and the recent and thereby transport themselves from flower to flower review by Gupta and Karmakar [51] provide valuable where they feed on pollen or nectar or other arthropods information on phytoseiid diversity. Till now, 217 species of [35]. An undescribed species of Proctolaelaps is predaceous Phytoseiidae have been described from India [47]. on phoretic nematodes during transport of both nematode and mite on the adult of a weevil species [36]. Melicharids Lifestyles of phytoseiid mites have not been investigated seriously in India. McMurtry et al. [52] classified phytoseiid mites based on their lifestyles: specialized predators of Tetranychus species (subtype I-a); specialized predators of web-nest-producing Family Otopheidomenidae mites (subtype I-b); specialized predators of tydeoids (subtype I-c); selective predators of tetranychid mites In this family, all the active stages are parasites of insects, (type II); generalist predators living on pubescent including many agricultural pests, throughout the world. (subtype III-a); generalist predators living on glabrous From India, Banerjee and Datta [37] reported suppression leaves (subtype III-b); generalist predators living in confined of the red bug, Dysdercus cingulatus (Fabricius), by an spaces on dicotyledonous plants (subtype III-c); generalist otopheidomenid mite, Hemipteroseius indicus (Krantz & predators living in confined spaces on monocotyledonous

http://www.cabi.org/cabreviews 4 CAB Reviews plants (subtype III-d); generalist predators from soil/litter Commercial utilization habitats (subtype III-e); and pollen-feeding generalist The phytoseiids known to keep their prey under predators (type IV). Besides, two additional lifestyles, viz. control and largely used in biocontrol programmes phytoseiids living on aquatic plants and phytoseiids capable are: A. andersoni, (Muma), A. swirskii, of piercing cells, were also proposed. (Oudemans), E. gossypii (El-Badry), E. hibisci, floridanus (Muma), G. occidentalis, degenerans (Berlese), arboreus Food habits (Chant), N. californicus, N. cucumeris (Oudemans), Phytoseiids are not generalist predators. Some phytoseiids N. fallacis, N. longispinosus (Evans), Phytoseiulus macropilis are carnivorous while a few prefer plant-derived food such (Banks), P. persimilis, S. newsami (Evans), Typhlodromus as pollen and nectar. Sometimes cannibalism and intraguild aripo (De Leon), T. caudiglans (Schuster), T. pyri and predation is also observed [53]. Phytoseiulus persimilis T. rickeri (Chant). prefers species that live in colonies, mostly Tetranychus species; Neoseiulus fallacis (Garman) feeds on Tetranychus Some successful biological control programmes with phytoseiids urticae (Koch) but not on Panonychus ulmi (Koch); Utilizing type I predator P. persimilis for control of Galendromus occidenatalis (Nesbitt) prefers strong-webbing Tetranychus spp. in greenhouse cucumber and outdoor phytophagous mites; (Chant) prefers to feed , type-II predator G. occidentalis for control of on species that are generally distributed; and Typhlodromus Tetranychus mcdanieli (McGregor) on apple, as well as using pyri (Scheuten) extracts plant juice. Various kinds of organophosphorus-resistant strain of G. occidentalis for fungi, including powdery mildews, may also serve as food control of Tetranychus pacificus (McGregor) on almond [e.g., aberrans (Oudemans)]. Phytoseiids are some of the examples of successful biocontrol usually cut into the cuticle of the prey by their chelicerae, programmes [55]. Periodical release of P. persimilis against inject a proteolytic enzyme, which liquefies its contents T. urticae in greenhouses was found successful [58]. and then the same is taken up by the predator [54]. They Phytoseiulus longipes (Evans) was found potentially useful also feed on the content of pollen by rupturing it with for control of Tetranychus evansi (Baker & Pritchard) [59]. chelicerae. Some phytoseiids are known to cut silken Type III predators Neoseiulus baraki (Athias-Henriot) and thread of mites by their chelicerae and thereafter Neoseiulus paspalivorus (De Leon) were found quite enter into their colonies (e.g., N. fallacis, P. persimilis) [55]. promising for control of coconut eriophyid mite, The food eaten by phytoseiids can be analysed by guerreronis (Keifer) [60]. Type III predator A. largoensis is electrophoresis of the gut contents and some such constantly associated with (Hirst) on coconut studies have been reviewed by Solomon et al. [56]. and it seems to be a potential predator of this mite [61]. However, so far, no phytoseiid mite has been shown as a specialist predator of tenuipalpid mites [62]. Alternative food Alternative food can be of importance for two reasons: Classical biological control First, it may help the predator in maintaining itself in The most successful classical biological control pro- a locality where or when the main prey population is low. grammes with phytoseiid mites are against the cassava Second, it helps in laboratory rearing of predators. Some green mite, Mononychellus tanajoa (Bondar), in Africa, species can multiply much faster on eriophyoids than on and and avocado mite in , USA. The other phytophagous mites [e.g., G. occidentalis on Aculus former was controlled by type III lifestyle species schlechtendali (Nalepa)]. Some can be reared effectively on Amblydromella manihoti (Moraes) and T. aripo, and that pollen [e.g., (Chant) on pollen of Vicia reduced the pest population by over 50%. In the case faba L.]. Even artificial foods like sugar solution and honey of European red mite on citrus [Panonychus citri can provide good nutrients. Typhlodromus pyri can survive (McGregor)] and avocado mite [Oligonychus punicae on apple leaf surface by feeding on plant sap [42]. (Hirst)], the predators of types II, III and IV were field-released, but only type IV predator Prey-searching ability (Athias-Henriot) spread extensively from the release site. A Kairomones play an important role in prey-searching ability pesticide-resistant strain of G. occidentalis was introduced by phytoseiids, either by contact or responding to plant into Australia against T. urticae on apple [63]. volatile compounds. Some of the volatile compounds are prey-specific [57]. Indian work on diverse aspects The phytoseiid species on which diverse types of work have been done in India are as below: Mass multiplication McMurtry et al. [55] discussed different techniques along (1) Amblyseius channabasavannai (Gupta & Daniel): Daniel with mentioning the sophisticated technique of Koppert®, [64] studied life cycle on R. indica and reported and for details the same may be referred to. developmental period of different stages.

http://www.cabi.org/cabreviews Salil Kumar Gupta and Prakya Sreerama Kumar 5 (2) Euseius alstoniae (Gupta): Dhooria [65] and Jose et al. of T. urticae [75]. Due to the inability of this mite to [66] studied predatory behaviour and rate of con- penetrate the dense web, having long generation time, poor sumption of food. searching ability as well as poor field population, its (3) Euseius coccosocius (Ghai & Menon): Its biology efficiency as a biocontrol agent is doubtful. This mite has been studied [38]. is also known to feed upon thrips, leafhoppers, aphids, (4) (Chant): Jagadish et al. [67] studied etc. Wallace [76] reported Anystis salicinus (L.) as a good life cycle at different temperatures. predator of red-legged earth mite, Halotydeus destructor (5) Euseius delhiensis (Narayanan & Kaur): Somchoudhary (Tucker), and the collembolan, Sminthurus viridis (L.), in [68] studied population fluctuation, while Australia. Betelvine pests such as Aleurocanthus rugosa Nageshchandra et al. [69] studied its biology. (Singh), Dialeurodes pallida (Singh), Tricentrus gibbosulus (6) Euseius finlandicus: A good amount of work on feed- (Walker), Membrothrips indicus (Hood), Zaniothrips ing potentiality, life cycle at different temperatures ricini (Bhatti), Mymarothrips garuda (Ramakrishna & and predator–prey interaction have been studied as Margabandhu) and Aroidothrips longistylus mentioned in Gupta [38]. (Ananthakrishnan) are known to be fed by anystid mites (7) (Evans): Its efficacy against the broad in West Bengal, India [77]. Walzia indiana (Smith-Meyer & mite, Polyphagotarsonemus latus (Banks), was studied Ueckermann) was found to feed upon eggs of coccids and by Hariyappa and Kulkarni [70]. aleurodes infesting betelvine [73]. Mathur [78] reported (8) Euseius rhododendronis (Gupta): Its predatory potenti- Anystis sp. as important predator of aphids of agricultural ality was worked out by Krishnamoorthy [71]. importance. Halliday and Paull [79] evaluated Chaussieria (9) Neoseiulus longispinosus: Its life cycle, food preference, capensis (Meyer & Ryke) as predator of H. destructor. Holm feeding potentiality, predator–prey interaction, mass and Wallace [80] evaluated A. baccarum in laboratory as a multiplication, effect of pesticide application, etc., have potential predator of cattle , microplus been worked out by different workers and all those (Canestrini). According to Hoy [81], anystid mites are have been reviewed in Gupta [38]. predators of limited value in biological control. (10) Phytoseiulus persimilis (Imported culture): Krishnamoorthy [4, 72] worked out its mass pro- Family Arrenuridae duction as well as life cycle. (11) Transeius tetranychivorus (Gupta): Diverse aspects of Members of this family occur in all types of freshwater this species have been studied [38]. habitats throughout the world. Some species are parasites of nematocerous Diptera or Odonata [82]. Many species Though A. largoensis and Scapulaseius suknaensis (Gupta) feed on ostracods and other invertebrates [83]. Larvae also appear to be quite promising, they have largely been of these aquatic mites are important biocontrol agents neglected. of mosquito larvae and adults. When mosquitoes come to the water surface for egg-laying, the mites get detached to them and further develop in water. Parasitization is Order Trombidiformes more on young larvae than on older larvae of mosquitoes. Parasitized mosquitoes lay fewer eggs [84]. Suborder Prostigmata Sharma and Prasad [85] and Bhattacharyya et al. [86] reported Arrenurus sp. parasitizing Culex mosquitoes in Family Anystidae India. Rajendran and Prasad [87] recorded Arrenurus madaraszi (Daday) parasitizing Anopheles mosquitoes and These exclusive predators are quite large, brightly coloured found it to be a good biocontrol agent. Sarkar et al. [88], and occur mostly on the undersurface of leaves but Biswas et al. [89] and Rahman et al. [90] were the other may shift to upper surface whenever disturbed (S.K. workers who reported Arrrenurus sp. as good parasitizing Gupta, personal observation). They are fast runners, and agent of anopheline mosquitoes. Malhotra and Mahanta if disturbed, they start making a whirling movement and [91] published a checklist of the mosquitoes parasitized by then drop down to the ground and escape (S.K. Gupta, Arrenurus in India. Mohan and Raghunatha Rao [92] personal observation). Anystidae is represented in India reported infestation of Arrenurus mite on aquatic insects. by ten species in three genera [73]. The most important They also highlighted the parasitizing effect of Arrenurus sp. as well as common species in many parts of the world, on Anopheles annularis (van der Wulp). including India, is Anystis baccarum (L.), which attacks tetranychid and tenuipalpid mites with agility. It holds the Family prey with forelegs and starts sucking from the body immediately. It is believed that it injects some toxin into Globally, this family is known by 135 species in 16 genera the prey body for paralyzing it before sucking [73]. Anystis and five subfamilies [83], but only 15 species are known agilis (Banks) is an important predator in citrus and apple from India [73, 93]. These free-living predators are quite orchards [74]. The adult female consumed 675.2 individuals large, mostly reddish or yellowish, with gnathosoma

http://www.cabi.org/cabreviews 6 CAB Reviews protruding anteriorly in the form of a snout, hence called observation). Cheyletus malaccensis is also known as a snout mites [83]. good predator of eggs of Rhyzopertha sp. and Trogoderma They are found under tree bark, on foliage, in soil sp. as well as eggs of Schizotetranychus sp. on paddy in and litter, and on algae. Many are important regulators of West Bengal and Brevipalpus sp. in Meghalaya [73]. Acaropsis phytophagous mites [84]. Bdellids can secrete a silken sollers (Rohdendorf) has been reported as an important thread to fasten their prey to the substrate, and are also biocontrol agent in storage system [103]. known to construct a moulting chamber [94]. Some feed on eggs and also on nematodes [95]. They are Family Cunaxidae generalized predators of spider mites, including species like the polyphagous T. urticae, praetiosa (Koch) on fruit These are fairly large, reddish/yellowish/brownish mites trees and Petrobia latens (Muller) on wheat. Bdella depressa mostly occurring on undersurface of leaves in the angles (Ewing) is a predator of B. praetiosa [96], and springtails. formed by veins (84; S.K. Gupta, personal observation). lapidaria (Kramer) efficiently fed on S. viridis,an They are very active predators capable of hunting the prey important pest of pasture in Australia and South Africa [97, by fastening it with a silken thread produced from their 98]. Although many undetermined species of this family mouth parts [83, 84]. As soon as disturbed, they jump and were reported from India in association with tetranychid try to escape (S.K. Gupta, personal observation). Smiley or tenuipalpid mites, actual feeding under field conditions [104] reviewed the world Cunaxidae wherein he included was not noticed [73]. 166 species. In India, there are 21 species in five genera [73]. The cunaxids are of two types as far as predatory Family behaviour is concerned [105]. The members of the subfamily Cunaxinae (e.g., species of Armascirus, Cunaxa These mites are free-living excepting a few that are parasitic and Dactyloscirus) are ambush predators having elongated to birds and mammals [84]. They are of various colours, palpi and long prodorsal trichobothria. They pounce upon slow-moving, mostly roundish and occur on leaves, twigs, as the prey and grab it with the silken thread secreted. The well as in several other habitats like stored products, house other group in the subfamily Coleoscirinae (e.g., species of dust, bird nests, etc. [84]. In all cases, they are very efficient Neoscirula and Coleoscirus) are cruise predators that search predators not only of the mites but also of immature stages out preys [83, 105]. Cunaxids are inhabitants on the bark of of insects (e.g., Coleoptera grubs) in stored products fruit trees infested by spider mites in the USA, but their where they are found abundantly (S.K. Gupta, personal effect on prey population is unknown [83]. Cunaxid mites observation). Cheyletidae is known by over 370 species in are known to be good predators of oystershell scale, 74 genera [83]. Plant-associated cheyletid mites are known Lepidosaphes ulmi (L.), on apple in North America [106]. in India by 15 species and ten genera [73]. The species that is often encountered in India is Cunaxa Hemicheyletia bakeri (Ehara) is known to suppress spider setirostris (Hermann), which feeds on Eutetranychus orientalis mites on citrus [99] and [100]. Due to the (Klein) on citrus in Punjab, tetranychids in Sikkim [73], poor searching ability, poor numerical and functional Oligonychus iseilemae (Hirst) on coconut in Kerala and responses to increasing population, H. bakeri Oligonychus mangiferus (Rahman & Sapra) on mango [73]. does not appear to be capable of suppressing the prey mite Cunaxa capreolus (Berlese) is another important predatory population in orchards. Often, it shows cannibalism. Muma species known to devour T. urticae, E. orientalis, etc. [73]. [101] reported it as a good predator of citrus mites in Florida. Mexecheles hawaiiensis (Baker) has also been Family reported as most abundant predator of (Geijskes) in Hawaii [102]. These are large-sized mites, variously coloured, with dense The common Indian species, Cheletogenes ornatus growth of hair on dorsal surface of body [84]. Though the (Canestrini & Fanzago), is known to feed on mango bud larval stages are parasitic to invertebrates, the post-larval mite, Aceria mangiferae (Sayed), and ber gall mite, Aceria stages are free-living predators of immature insects cernuus (Massee). It is believed that, if properly utilized, and other mites [84]. Lasioerythraeus whitcombi (Smiley) C. ornatus can help in controlling these two eriophyoid pests is known to feed on eggs of cotton bollworm in the [73]. Parachyletia pyreformes (Banks) has been reported to USA [107]. Larvae of Erythrites jacksoni (Southcott) were be a good predator of grapevine mite in Kashmir valley [73]. reported to parasitize the brachypterous females of grass- Cheyletid mites frequently encountered in storage are hopper, Brachaspis collinus (Hutton) in New Zealand [108]. Cheyletus eruditus (Schrank), Cheyletus malaccensis Species of Erythraeus are known to parasitize some (Oudemans) and Cheyletus fortis (Oudemans) (73; S.K. hemipterans [109]. Gupta, personal observation). They readily attack acarid, Balaustium putmani (Smiley) fed on 20 eggs of P. citri, glycyphagid and pyroglyphid mites as well as grubs of while nymphs consumed 89–100 eggs [110]. Besides mites, beetles such as Sitophilus oryzae L. and Tribolium castaneum it also is known to feed upon scale insects, moths and (Herbst), thus acting as a good limiting factor in reducing their eggs. However, it cannot tolerate pesticide application stored product pests (73; S.K. Gupta, personal in orchards [111]. Balaustium murarum (Hermann) is a

http://www.cabi.org/cabreviews Salil Kumar Gupta and Prakya Sreerama Kumar 7 good egg predator of apple torricid, Zeiraphera diniana australis (Say) [62]. Some like Hydryphantes tenuabilis (Guenee) [112]. (Marshall) feed on dragonfly eggs in laboratory rearing In India, 18 species have been reported to be associated [131]. Larvae of Wandesia thermalis (K. Viets) parasitize with plants [73, 113]. Balaustium sp. was reported to brine flies and adult insects feed on eggs of these flies [132]. consume Tetranychus sp. on brinjal in West Bengal [73]. Ghai and Ahmed [114] reported Bochartia sp. parasitizing Family Iolinidae leafhoppers. Seshu Reddy and Davies [115] reported Abrolophus sp. parasitizing sorghum shoot fly, Atherigona The family Iolinidae, which includes genera like soccata (Rondani), in Hyderabad. Sorghum shoot bug, Pronematus and Proctotydeus, consists of 125 described Peregrinus maidis (Ashmead), was reported to be parasitized species under 36 genera [83]. These are free-living and by Erythraeus sp. in Dharwad [116]. The soapberry bug, occur in soil, bird nests, plants, stored food products, etc. Leptocoris augur (Fabricius), is also parasitized by Leptus sp. (S.K. Gupta, personal observation). Pronematus elongatus [117, 118]. Agarwal and Dhiman [119] reported Leptus sp. (Baker) is known to feed on O. mangiferus on grapevine in parasitizing Cletus signatus (Walker) in Uttar Pradesh. Kashmir, India [133]. Pronematus fleschneri (Baker) appears Rawat [120] reported Bochartia sp. feeding on various to be a good predator on eggs of Tetranychus neocaledonicus phytophagous hemipterans in Gwalior. Rawat and Saxena (André) on brinjal as well as on the citrus mite, E. orientalis [121] reported Bochartia sp. feeding upon nymphs and [73]. Pronematus sextoni (Baker) also has been reported adults of sorghum shoot bug, P. maidis, and the parasitiza- to feed upon E. orientalis [134]. Pronematus ubiquitus feeds tion was 3.7–15.5%. Helopeltis antonii (Signoret) was also on fig eriophyid in California and E. orientalis in Egypt reported to be paratisized by Bochartia sp. [122]. Tandon [62]. Pronematus sp. fed on B. phoenicis on tea in India and Lal [123] recorded an unidentified species of [62]. Homeopronematus anconai (Baker) is reported to feed Erythraeidae parasitizing Drosicha mangiferae (Green). upon russet mite, lycopersici (Massee), in California, USA, reducing the population substantially Family Hydrachnidae [135]. Species of Proctotydeus are insect associates and are found in galleries [136]. Hydrachinidae include many species of Hydrachna that inhabit temporary ponds, pools and lakes [124]. Family Pionidae Adults of Hydrachna conjecta (Koenike) feed on eggs of water boatman species, Sigara striata (L.), laid in submerged This family is represented in the world by many species aquatic plants [125]. Larval Hydrachna are parasites of many under seven subfamilies [83]. They inhabit water bodies aquatic hemipteran and coleopteran insects [126]. Mohan [137]. The larval forms of this family are known to parasitize [127] reported a Hydrachna sp. parasitizing the aquatic Chironomidae [128]. These colourful aquatic mites are bug Lethocerus indicus (Lepeletier & Serville) in the river voracious feeders of mosquito larvae [e.g., Piona nodata Bhilangana in Tehri Garhwal Himalaya in India. (Müller)] [84]. Piona species are known to feed upon small aquatic arthropods like cladocerans [138]. However, this Family Hydryphantidae group has not been properly explored for its utility in bio- control [129]. Only six species are known from India [139]. Across the world, this family is known by several species included in 12 subfamilies. These colourful, soft-bodied, Family Pterygosomatidae water mites usually occur in ponds and temporary pools. The larvae are parasitic, and deutonymphs and adults are These red, small to medium size mites are good predators predatory; protonymphs and tritonymphs remain quies- of reduviid bugs and cockroaches [83, 140]. Pimeliaphilus cent. They feed on mosquitoes, biting midges, black flies plunifer (Newell & Ryckman) is known to parasitize blood- and horse flies [62] of medical and veterinary importance. sucking bugs of Triatomidae [140]. The mortality may not Larvae of Wandesia species are parasites of stoneflies be because of sucking of body fluid but may be due to [128]. Euthyadine larvae parasitize aquatic Hemiptera, injection of a toxin during feeding [141]. This family is Diptera and Trichoptera, and opportunistically use terres- known by about 120 species. Most of the described species trial hosts belonging to Collembola, Thysanoptera and are parasites of lizards, tortoises or arthropods in Africa, some Hemiptera that sometimes occur on wet vegetation Asia, Australia and America [83]. They feed on body fluid of [83]. The larvae of some hydryphantid species parasitize their hosts and sometimes cause severe disorders like mosquito larvae, but the post-larval stages are free-living anaemia and intense skin irritation [142]. predators consuming mosquito eggs [129]. Thyas sp. feeds on mosquito eggs [130]. The pattern of mosquito parasit- Family Pyemotidae ism differs from that of members of other water mite families because they attack hosts when returning to water As many as 20 species in this family have been identified, bodies not as these adults first emerged [62]. They also all under one genus, Pyemotes. The members of this family act as natural enemies of the water measurer, Hydrometra are mainly parasites of insects in the orders Thysanoptera,

http://www.cabi.org/cabreviews 8 CAB Reviews Hemiptera, , Coleoptera, Diptera and reddish, mostly occur on undersurface of leaves, and are Hymenoptera [83]. These are milky white, fast-moving slow in their movement unlike phytoseiid mites, which run mites and are frequently found in stored-product habitats, very fast when chased. This family is represented from the straw, etc., parasitizing some injurious insects. It is also world by about 300 species in 30 genera [83]. known to kill various coleopteran and lepidopteran insects The important genera which are known to feed in stored products [84]. The straw itch mite, upon spider mites are Agistemus, Eryngiopus, Stigmaeus (Lagreze-Fossat & Montane), is an effective biocontrol and Zetzellia [62, 81]. Laing and Knop [155] studied agent of many insect pests. It attacks all stages of Plodia the potential use of stigmaeids in biocontrol programmes. interpunctella (Hübner) (Lepidoptera). It was introduced Agistemus species prey upon spider mites and other for control of fire ant, Solenopsis invicta (Buren), in the orchard-associated arthropods in various parts of the USA [143]. Moser et al. [144] reported two groups of world [96, 156]. Agistemus fleschneri (Summers) is a pyemotid species, the ones with poison and the other predator of the tetranychids, P. ulmi, Tetranychus cinnabar- without. inus (Boisduval) and Tetranychus kanzawai (Yokoyama). It In India, Singh and Singh [145] reported that Pyemotes feeds upon P. ulmi and T. urticae in orchards in the USA and herfsi (Oudemans) parasitized diapausing pink bollworm, Egypt, respectively, and the tenuipalpid mite B. phoenicis gossypiella (Saunders), on cotton. This mite on tea in Indonesia. Agistemus floridanus is a predator of has been used in biological control of P. gossypiella in Eotetranychus sexmaculatus (Riley) and various scale Egypt [146]. Nath and Balan [147] reported it on the insects in Florida, USA. Agistemus exsertus (Gonzalez- same pest in Kashmir. Dhiman and Mittal [148] recorded Rodriguez) is a predator of P. citri [81, 157]. Agistemus Pyemotes sp. parasitizing the common Indian housefly, faneri (Dosse) is a predator of tetranychids, whereas M. domestica. Dakshinamurthy et al. [149] reported A. longisetus is a predator of P. ulmi [81]. Agistemus exsertus Pyemotes ventricosus (Newport) parasitizing Sitotroga cerea- devoured 60 eggs or 40 immatures of the eriophyid lella (Olivier). A. lycopersici in a day [158]. Agistemus fleschneri is another Species of Pyemotes may attack humans, especially good predatory mite known to feed on eggs of apple granary workers and food-handlers, and domestic mites, consuming over 430 eggs or 1498 larvae during its , when they are exposed to infested grains. They lifetime [84]. also cause other symptoms like fever and lymphadenopathy Zetzellia mali (Ewing) is an important natural enemy [150]. The resultant bites can induce dermatitis (often of apple mites in North America and [62]. White called ‘grain itch’ or ‘hay itch’), a factor that seriously and Laing [159, 160] worked out the intrinsic rate hinders the use of these mites for biocontrol [62]. of increase and mean generation time of Z. mali that fed upon A. schlechtendali on apple. It has preference for Family Raphignathidae A. schlechtendali over P. ulmi eggs in apple orchards in Canada [161]. Zetzellia mali also feeds upon eggs of the This family is known from the world by over 40 species phytoseiids G. occidentalis and T. pyri [162]. [83], but only four species are known from India [73, 151]. India has 27 stigmaeid species in six genera [73, 151, 163]. A raphignathid was found to feed on tetranychid Agistemus fleschneri is one of the foremost species that eggs on leaves of some medicinal plants (S.K. Gupta, shows good biocontrol potential in India [73, 164, 165]. personal observation). Raphignathus youngi (Hirst) was Its prey species are: Brevipalpus obovatus (Donnadieu) on reported to be associated with phlebotomid sandflies in guava in West Bengal; tetranychids in Delhi; A. mangiferae Surat [152]. on mango in Manipur; and T. urticae on okra in Assam [73]. Agistemus herbarius (Kuznetzov & Wainstein) has Family been reported to feed upon O. mangiferus and T. urticae on grapevine in Kashmir [73, 133]. Agistemus industani These are moderately large, active predators of small (Gonzalez-Rodriguez) is a very important stigmaeid that is arthropods and are globally distributed [83]. Many of abundantly available and also has good predatory potential. the members of this family have been found on a range of Rai and Singh [166] studied its feeding behaviour on insect hosts, including scale insects [153]. Neophanolophus Tetranychus ludeni (Zacher) on mulberry in Varanasi sp. was reported to occur on Erythroneura leafhopper in and reported its voracious feeding on eggs. The feeding India [154]. potential of the female was three times higher than that of the male. Agistemus macrommatus (Gonzalez-Rodriguez) Family Stigmaeidae fed upon T. ludeni on cowpea in Uttar Pradesh and T. neocaledonicus on Calotropis in Lakshadweep [73]. Among the predatory mites that are of potential Agistemus terminalis (Quayle) fed upon Brevipalpus in importance, the members of Stigmaeidae are certainly Sikkim and Acaphylla theae (Watt) on tea [73]. Eryngiopus worthy to be reckoned. These are considered next only to coimbatorensis (Gupta & David) was reported to totally Phytoseiidae as regards their efficiency as biocontrol agents eliminate the sugarcane scale, Melanaspis glomerata as observed by the present authors. These are yellowish or (Green) [167].

http://www.cabi.org/cabreviews Salil Kumar Gupta and Prakya Sreerama Kumar 9 Family favoured and predated on the eggs, but not on the adults of T. urticae. This family is known from the world by 530 species in Species of Dinothrombium are among the largest (meas- 40 genera [83]. These fast-moving mites are light yellowish uring up to 12 mm) of known mites that inhabit deserts in or blackish or light greenish. The male often shows a typical western USA and their mass emergence occurs following pre-copulation behaviour of carrying an adult pharate heavy summer rains giving a spectacular velvety appearance female on its posterior dorsum using the caudal genital to a parched desert floor [180]. Dinothrombium pandorae papillae and sometimes with its modified leg IV to hold it (Newell & Tevis) is parasitic to grasshoppers. Eutrombidium [168, 169] so that it could mate as soon as the adult female larvae parasitize grasshoppers causing 75% . emerges. This family shows the greatest diversity in feeding Infestation of this mite hinders the flight capability of its habits of all acarine families [169]. Some are associated insect hosts. Podothrombium larvae are parasitic to aphids with arthropods as predators, parasites or phoretics; many [181]. are fungivores, some species are phytophagous [a few, such In India, Chandra [182] reported that Allothrombium sp. as P. latus, or cyclamen mite, Phytonemus pallidus (Banks), suppressed the buzzing grasshopper [Sphingonotus savignyi are important crop pests]; and there are others whose (Saussure)] population. Dhiman and Dhiman [183] re- lifestyles are still undetermined [169]. Tarsonmenids corded Microtrombidium sp. as a parasite of M. domestica. are known to feed on eggs of tetranychid and tenuipalpid Dhiman et al. [184] recorded Microtrombidium saharanpuri mites [170]. The larvae and adults of Acaronemus (Dhiman) as a parasite of the green bottle fly, Lucilia caesar prey upon tetranychid and tenuipalpid mites on foliage (L.). Mittal and Dhiman [185] studied the population dy- [83]. Some like Dendroptus prey upon gall-producing namics of M. saharanpuri, an ectoparasite of Indian housefly eriophyoids or on eggs of tydeid mites. The larvae and the same authors [186] studied distribution of that of Acarapis are parasites of adult honey bees. Iponemus species. Suhas and Jai Rao [187] reported Trichotrombidium sp. parasitizes pestiferous bark beetles of Ips. Acaronemus muscarum (Kolonev) parasitizing the housefly in India. destructor (Smiley & Landwehr) feeds on eggs of phytophagous mites [84]. The members of the genera Family Pseudotarsonemoides, Ununguitarsonemus, Heterotarsonemus and some Tarsonemus are phoretic, while members These are soft-bodied, weakly to highly sclerotized, green- of the genera Steneotarsonemus, Polyphagotarsonemus, ish yellow or pink mites having striations or reticulations Hemitarsonemus and Phytonemus are economically impor- on body. As many as 340 described species under 30 genera tant phytophagous mites. Some tarsonemids may serve are known from the world [83], and India represents only as supplementary diet for predaceous phytoseiids on eight species [73]. These mites are predators and can plants [40]. Some tarsonemids even cause human feed on fungi, pollen or even plants [62]. Some tydeids disorders like skin rash and have also been isolated from (e.g., Lorryia and species) are of phoretic nature on the surface of skin of a person having dermal abnormalities noctuid moths [188]. Tydeid mites occur in diverse habitats [171, 172]. They are considered to be one of the causal like moss, litter, straw, soil, humus, fungi, bird nests, stored agents of human dust allergy [173]. food products as well as on plants [189]. At least one species, Tydeus molestus (Moniez), has been reported to Family bite human beings and domestic animals [83]. Reports are available on their predatory habits [190] This is a large family, distributed worldwide and is as well as phytophagy [191, 192]. Some reports suggest represented by several species under 20 genera in three its feeding on nematodes and eriophyids [81]. Tydeus subfamilies [174]. The larvae of these large, velvety mites californicus (Banks) was reported to feed on citrus bud are parasitic to invertebrates whereas the post-larval stages mite, Aceria sheldoni (Ewing) [81]. Knop and Hoy [193] are free-living. The larvae of species parasitize reported tydeid predators in vineyards. Those tydeids many insects and . which feed on pollen, fungi and plant tissues live long and Allothrobium lerouxi (Moss) feeds on aphids and cater- produce more progeny [62]. They also prey on spider mites pillars in apple orchards [175]. Allothrombium pulvinum and eggs of citrus flat mites [62]. Tydeus caudatus (Dugès) (Ewing) is an ectoparasite on aphids whereas its deuto- feeds on grapevine downy mildew [81]. nymphs and adults are predatory towards aphids and The association of tydeids with spider mites does not eggs of lepidopterans; the protonymphs and tritonymphs always signify their feeding on the latter, though their are quiescent. It is an important predator of spider potential as biocontrol agents cannot be ruled out. mites in peach orchards [176] and cotton fields [177] However, because of their low field populations, poor in China. According to Chen et al. [178], A. pulvinum is a searching ability and low feeding capacity, they may not have major limiting factor of cotton aphid, Aphis gossypii the desired influence on the spider mite population. (Glover), and can successfully suppress its population. Further, studies relating to their biological aspects along Zhang [179] studied the functional response of A. pulvinum with numerical and functional responses may throw new deutonymphs on T. urticae. The deutonymphs light on their biocontrol utility.

http://www.cabi.org/cabreviews 10 CAB Reviews Order Sarcoptiformes Because of non-specificity, their predatory efficiency is variable. For example, Luck et al. [205] reported that Suborder Oribatida Hemisarcoptes cooremani (Thomas) was not a good candi- date for augmentative release against California red scale, Cohort Astigmatina Aonidiella aurantii (Maskell). Scales attacked by 5–10 mites do not produce progeny and those attacked by more than Family ten mites and fed would die [206].

Members of the Acaridae belong to diverse ecological habitats. Some acarids are known to act as predators of phytophagous mites, nematodes and rootworms. Others Suborder Oribatida are phoretic, nidicolous, pollen-feeders or plant tissue- feeders. However, generally, they are serious pests of Cohort excluding Astigmatina stored products (e.g., siro L.) [194]. As many as 500 species in 80 genera are known from the This suborder is represented from the world by over 9000 world [195]. species in 172 families [207]. From India, over 662 species, Though the majority of acarids are harmful pests of 42 subspecies, 88 families, 244 genera and 103 subgenera stored products, some may be beneficial [81]. are known [208]. These are primarily found in the soil and putrescentiae (Schrank) is known to effectively feed on corn litter, others are arboreal [81]. rootworm [Diabotrica undecimpunctata howardi (Barber)]. It Oribatid mites constitute almost 70% of soil micro- is known to prefer this prey over [29]. This species arthropod fauna. Soil oribatids bring about diverse activities has also been reported to feed on dead and living adults and like soil aeration, mixing soil and microbes, assisting in eggs of the grapevine mite, Daktulosphaira vitifolii (Fitch) nutrient recycling, helping in decomposition of soil [196]. voraciously feeds on nema- organic matter and enrichment of soil fertility, helping in todes in laboratory culture [197]. It has also been reported mineralizing plant nutrients and stimulating species succes- to feed upon Meloidogyne javanica (Treub), a serious plant sion; some even act as predators [81]. Oribatids have parasitic nematode. different types of feeding behaviour. The largest species Species of inhabit bulbs and Caloglyphus feed on leaf litter whereas the smallest species feed on species inhabit compost and decaying fungi [198]. Members fungi, and other microbes. They may be micro- of Hoogstraalacarus live on gills of hermit crab [199]. Kuzinia phytophages, panphytophages, zoophages, necrophages species are obligatory associates of bumblebees and feed or coprophages. The panphytophages depend upon the on pollen [200]. Caloglyphus sp. was reported to infest egg pollen algae, lichens, decomposed humus fungal hyphae, pods of locust and could effectively control it. In India, etc. [81]. Some can be reared on nematodes. Rearing is also sp. was reported to feed upon sugarcane white possible on mushrooms, chopped lichens, decomposed grub [73]. leaves or on artificial diets like dextrose and casein or brewer’s yeast [81]. Some oribatids are pests of crops and Family intermediate hosts of cestode parasites [209]. At least 47 species of oribatid mites under 32 genera transmit It is one of the most diverse groups of Astigmatina. Some 12 species of cestodes [210]. are common inhabitants of house dust and stored food Xylobates seminudus (Hammer) is known to devour products [e.g., Nanacarus minutus (Oudemans)] and some nematodes in India. An individual mite would require – are associates of cortical and subcortical insects [201]. 35 45 min for completion of feeding activity [211]. Species of Hemisarcoptes have parasitic habits. Some have phoretic association with beetles [81]. This family is known by 31 species under 11 genera from the world [195]. Indian Summary and Conclusion information on the diversity of this family is poor. These whitish, soft-bodied mites are also known as predators The global number of extant mite species is believed to vary of armoured scale insects, attacking both adults and eggs, from 0.5 to 1 million [212] and a substantial figure could and therefore, are regarded as good agents for managing be predatory, and possibly, parasitic mites. India is still scale insects [62, 202]. Hemisarcoptes malus (Shimer) lagging behind several countries in terms of unearthing causes mortality of L. ulmi, on apple in eastern Canada new species of mites in general and predatory species in [203]. Hemisarcoptes coccophagus (Meyer) is reported to particular. In view of the diversity of predatory and parasitic feed on date palm scale insect, Parlatoria blanchardi mites in India, introduction of exotic species does not (Targioni-Tozzetti), in Africa [204]. When it fed on appear to be sustainable or economically sound. Efforts Lepidosaphes beckii (Newman) it turned purple, but it was should be directed at conservation and profitable utilization red when fed on Epidiaspis leperii (Signoret) and yellow of the indigenous species not only against the pests of when fed on Quadraspidiotus juglansregiae (Comstock) [81]. agri-horticultural crops, but also against pests of medical

http://www.cabi.org/cabreviews Salil Kumar Gupta and Prakya Sreerama Kumar 11 and veterinary importance. Some of the species with References potential biocontrol value are: 1. van Lenteren JC, Bale J, Bigler F, Hokkanen HMT, (1) Against pests of agri-horticultural crops and stored Loomans AJM. Assessing risks of releasing exotic biological grain: control agents of arthropod pests. Annual Review of – (i) Phytoseiidae – A. channabasavannai, A. herbicolus, Entomology 2006;51:609 34. A. largoensis, E. alstoniae, E. finlandicus, E. ovalis, 2. Rao VP, Datta B, Ramaseshiah G. Natural enemy complex N. fallacis, N. longispinosus, S. suknaensis, of flushworm and phytophagous mites on tea in India. T. tetranychivorus. Tea Board Scientific Publications Series (No. 5) 1970; 53 pp. (ii) Stigmaeidae – A. fleschneri, A. industani, E. 3. Sankaran T. Natural enemies introduced in recent years for coimbatorensis biological control of agricultural pests in India. Indian Journal – (iii) Anystidae – A. baccarum, W. indiana of Agricultural Sciences 1974;44:425 33. (iv) Cunaxidae – C. setirostris 4. Krishnamoorthy A. A simple method for mass rearing of (v) Iolinidae – P. fleschneri an exotic predacious phytoseiid mite, Phytoseiulus persimilis – (vi) Pyemotidae – P. tritici A.H. Journal of Biological Control 1988;2:53 55. (2) Against pests of medical, veterinary or public health 5. Singh SP. Fifteen years of AICRP on biological control. importance Technical Bulletin No. 8, Project Directorate of Biological (i) Arrenuridae – Arrenurus spp. (target Anopheles/ Control (ICAR), Bangalore, India; 1994. Culex mosquitoes) 6. Walter DE. Nematophagy by soil arthropods from the (ii) Trombidiidae – M. saharanpuri (target housefly and shortgrass steppe, Chihuahuan Desert and Rocky Mountains green bottle fly) of the central . Agriculture, Ecosystems & Environment 1988;24:307–16.

Our knowledge in India is lacking in the study of diversity 7. Walter DE, Halliday RB, Lindquist EE. A review of the genus and taxonomic aspects but equally so in bioecology, mass Asca (acarina: Ascidae) in Australia, with three descriptions of culture techniques and field utilization. Hence, it is strongly three new leaf-inhabiting species. Invertebrate Taxonomy – suggested to intensify studies in these areas. 1993;7:1327 47. The indigenous predators and parasites seriously 8. Lindquist EE, Walter DE. Antennoseius (Vitzthumia) janus need conservation through stopping indiscriminate and n. sp. (Acari, Ascidae), a mesostigmatic mite exhibiting unregulated use of pesticides in crops. Awareness needs adult female dimorphism. Canadian Journal of Zoology 1989;67:1291–310. to be generated among farmers on conservation and augmentative biocontrol options. Wherever possible, 9. Lindquist EE, Evans GO. Taxonomic concepts in the Ascidae, genetic manipulation may be attempted to produce with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the pesticide-resistant strains. Studies on food preference, Entomological Society of Canada 1965;47:1–64. alternative food, feeding potentiality, and functional and numerical responses of the potentially important predatory 10. Evans GO, Till WM. Mesostigmatic mites of Britain and Ireland (: Acari: ). An introduction to their species are also recommended. Finally, estimating the external morphology and classification. Transactions of the cost–benefit ratio of using predatory mites in lieu Zoological Society of London 1979;35:139–270. of adopting conventional chemical control methods is 11. Santos VVD, Tixier M-S. Integrative taxonomy approach for a must. analysing evolutionary history of the tribe Euseiini Chant & It is concluded, nonetheless, that indigenous predatory McMurtry (Acari: Phytoseiidae). Systematics and Biodiversity or parasitic mites deserve more attention and efforts than 2018;16:302–19. do the exotic species as past experiences with the latter 12. Walter DE, Lindquist EE. Life history and trophic behavior of were not all that successful (personal observations of mites in the genus Lasioseius (acari : Mesostigmata: Ascidae) authors). Furthermore, to reduce the perceived or actual from grassland soils in Colorado, with taxonomic notes and a risks of releasing exotic predatory mites for inundative or description of a new species. Canadian Journal of Zoology – augmentative biocontrol of indigenous pests, it is prudent 1989;67:2797 813. to first consider the use of native species [1]. 13. Evans GO. A revision of the British Aceosejinae (Acarina: Mesostigmata). Proceedings of the Zoological Society of London 1958;131:177–229. Acknowledgements 14. Chant DA. The subfamily Blattisocinae Garman (=Aceosejinae Evans) (Acarina: Blattisocidae Garman) (=Aceosejidae Baker The authors thank the undisclosed reviewer for and Wharton) in North America, with descriptions of new the constructive comments and additional inputs that species. Canadian Journal of Zoology 1963;41:243–305. greatly improved the manuscript. The kind invitation from 15. Halliday RB, Walter DE, Lindquist EE. Revision of the Dr David Hemming, Commissioning Editor, CAB Reviews, Australian Ascidae (Acarina: Mesostigmata). Invertebrate to write this review is gratefully acknowledged. The first Taxonomy 1998;12:1–54. author thanks the Secretary, Ramakrishna Mission, Kolkata, 16. Lindquist EE, Krantz GW, Walter DE. Order Mesostigmata. for the facilities provided for writing this review. In: Krantz GW, Walter DE, editors. A Manual of Acarology,

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